Slow frequency repetitive transcranial magnetic stimulation affects reaction times, but not priming effects, in a masked prime task.

OBJECTIVE: Slow frequency repetitive transcranial magnetic stimulation (rTMS) reduces motor cortex excitability, but it is unclear whether this has behavioural consequences in healthy subjects. METHODS: We examined the effects of 1 Hz rTMS (train of 20 min; stimulus intensity 80% of active motor threshold) over left motor or left premotor cortex on performance in a visually cued choice reaction time task, using a 'masked prime' paradigm to assess whether rTMS might affect more automatic motor processes. Twelve healthy volunteers participated. RESULTS: Motor cortex rTMS and, to a lesser extent, premotor cortex rTMS resulted in a slowing of right (stimulated) hand responses, but not of left (unstimulated) hand responses. In a control experiment, rTMS of the left somatosensory cortex did not lead to slower right hand responses. DISCUSSION: We conclude that long trains of low intensity 1 Hz rTMS over the motor or premotor cortex can have subtle behavioural consequences outlasting the stimulation. rTMS did not affect the modulation of reaction times by subliminal primes, suggesting that priming effects triggered by subliminal primes are not generated at the level of motor or pre-motor cortex.

Inhibition of subliminally primed responses is mediated by the caudate and thalamus: evidence from functional MRI and Huntington's disease.

Masked prime tasks have shown that sensory information that has not been consciously perceived can nevertheless trigger the preactivation of a motor response. Automatic inhibitory control processes prevent such response tendencies from interfering with behaviour. The present study investigated the possibility that these inhibitory control processes are mediated by a cortico-striatal-pallidal-thalamic pathway by using a masked prime task with Huntington's disease patients (Experiment 1) and with healthy volunteers in a functional MRI (fMRI) study (Experiment 2). In the masked prime task, clearly visible left- or right-pointing target arrows are preceded by briefly presented and subsequently masked prime arrows. Participants respond quickly with a left or right key-press to each target. Trials are either compatible (prime and target pointing in the same direction) or incompatible (prime and target pointing in different directions). Prior behavioural and electrophysiological results show that automatic inhibition of the initially primed response tendency is reflected in a 'negative compatibility effect' (faster reaction times for incompatible trials than for compatible trials), and is shown to consist of three distinct processes (prime activation, response inhibition and response conflict) occurring within 300 ms. Experiment 1 tested the hypothesis that lesions of the striatum would interrupt automatic inhibitory control by studying early-stage Huntington's disease patients. Findings supported the hypothesis: there was a bimodal distribution for patients, with one-third (choreic) showing disinhibition, manifested as an absent negative compatibility effect, and two-thirds (non-choreic) showing excessive inhibition, manifested as a significantly greater negative compatibility effect than that in controls. Experiment 2 used fMRI and a region of interest (ROI) template-based method to further test the hypothesis that structures of the striatal-pallidal-thalamic pathway mediate one or more of the processes of automatic inhibitory control. Neither prime activation nor response conflict significantly engaged any ROIs, but the response inhibition process led to significant modulation of both the caudate and thalamus. Taken together, these experiments indicate a causal role for the caudate nucleus and thalamus in automatic inhibitory motor control, and the results are consistent with performance of the task requiring both direct and indirect striatal-pallidal-thalamic pathways. The finding that Huntington's disease patients with greater chorea were disinhibited is consistent with the theory that chorea arises from selective degeneration of striatal projections to the lateral globus pallidus, while the exaggerated inhibitory effect for patients with little or no chorea may be due to additional degeneration of projections to the medial globus pallidus.

Learning to ignore the mask in texture segmentation tasks.

Although traditionally texture segmentation has been regarded as an automatic, preattentive process, participants confronted with texture segmentation in experimental settings (i.e., with brief presentation time and subsequent masking) are initially unable to perform the task. According to perceptual learning concepts, participants must learn to fine-tune their sensory channels before perception improves under restricted viewing conditions. The present article proposes an alternative perspective that emphasizes the role of the mask. Four experiments showed that the amount of observed learning depends on the structural and temporal homogeneity or heterogeneity of the mask. The authors suggest that learning consists of separating the task-relevant signal stemming from the texture from the task-irrelevant signal of the mask and of ignoring the mask.

Partial response activation to masked primes is not dependent on response readiness.

Masked primes presented foveally prior to a target trigger an initial partial activation of their corresponding response, followed by an inhibition of the same response. The latter phase results in performance costs on compatible trials and performance benefits on incompatible trials relative to neutral trials (negative compatibility effect). The present study investigated whether this activation-follow-by-inhibition process depends on the overall or specific state of response readiness. In two masked priming experiments, response readiness was manipulated by varying the relative frequency of Go-trials in a Go/NoGo task (Exp. 1) and the relative frequency of left- and right-hand responses in a 2-alternative choice reaction time task (Exp. 2). In both experiments, mean reaction times were longer for infrequent responses than for frequent responses. However, negative compatibility effects were not affected by response frequency. This result indicates that neither the general ability of masked primes to elicit a partial motor activation nor the specific time course of this process is dependent on response readiness. It is concluded that response readiness affects the execution of an overt response rather than the initial activation of this response.

Response facilitation and inhibition in manual, vocal, and oculomotor performance: evidence for a modality-unspecific mechanism.

In 2 experiments (N = 10, Experiment 1; N = 16, Experiment 2), the authors investigated whether evidence for response facilitation and subsequent inhibition elicited by masked prime stimuli can be observed for output modalities other than manual responding. Masked primes were followed by target stimuli that required a 2-choice manual, saccadic, or vocal response. Performance was measured for compatible trials in which primes and targets were identical and for incompatible trials in which they were mapped to opposite responses. When primes were presented centrally, performance benefits were obtained for incompatible trials; whereas for peripherally presented primes, performance benefits were found in compatible trials. That pattern of results was obtained for manual responses and for saccadic eye movements (Experiment 1), demonstrating that those effects are not mediated by specialized dorsal pathways involved in visuomanual control. An analogous pattern of effects was found when manual and vocal responses were compared (Experiment 2). Because vocal responding is controlled by the inferotemporal cortex, that result shows that prime-target compatibility effects are not primarily mediated by the dorsal stream, but appear to reflect modality-unspecific visuomotor links that allow rapid activation of motor responses that may later be subject to inhibition.

Chunking processes in the learning of event sequences: electrophysiological indicators.

The present study investigated whether effects of implicit learning (IL) are due to well-learned and explicitly represented parts of the stimulus material ("chunks"). To this purpose, event-related brain potentials (ERPs) were recorded during an oddball-version of a serial reaction time (RT) task: At unpredictable positions within a 16-item letter sequence, single deviant items replaced an item of the repeatedly presented standard sequence. After acquisition, the "process dissociation procedure" (Jacoby, 1991) was adopted to identify explicitly learned sequence parts for each participant. Acquisition of sequence knowledge was reflected in faster RTs for standard items than for deviant items and in enhanced N2b and P3b components for deviant items. While the ERP effects were obtained for explicitly represented sequence parts only, RT effects were independent of subsequent reproduction performance. These results indicated that (1) ERPs are a valid measure of explicit knowledge, (2) implicit and explicit knowledge coexist in serial RT tasks, and (3) chunking processes play a major role in the acquisition of explicit knowledge about event sequences.

A central-peripheral asymmetry in masked priming.

Masked primes presented prior to a target result in behavioral benefits on incompatible trials (in which the prime and the target are mapped onto opposite responses) when they appear at fixation, but in behavioral benefits on compatible trials (in which the prime and the target are mapped onto the same response) when appearing peripherally. In Experiment 1, the time course of this central-peripheral asymmetry (CPA) was investigated. For central primes, compatible-trial benefits at short stimulus onset asynchronies (SOAs) turned into incompatible-trial benefits at longer SOAs. For peripheral primes, compatible-trial benefits at short SOAs increased in size with longer SOAs. Experiment 2 showed that these effects also occur when primes and targets are physically dissimilar, ruling out an interpretation in terms of the perceptual properties of the stimulus material. In Experiments 3 and 4, the question was investigated as to whether the CPA is related to visual-spatial attention and/or retinal eccentricity per se. The results indicate that the CPA is independent of attentional factors but strongly related to the physiological inhomogeneity of the retina. It is argued that central and peripheral primes trigger an initial motor activation, which is inhibited only if primes are presented at retinal locations of sufficiently high perceptual sensitivity. The results are discussed in terms of an activation threshold model.

Effects of masked stimuli on motor activation: behavioral and electrophysiological evidence.

Three experiments investigated the influence of unperceived events on response activation. Masked primers were presented before a target. On compatible trials, primes and targets were identical; on incompatible trials, opposite responses were assigned to them. Forced-choice performance indicated that prime identification was prevented by the masking procedure, but overt performance and motor activation as mirrored by the lateralized readiness potential (LRP) were systematically influenced by the prime. The direction of these effects was unexpected: Performance costs for compatible and performance benefits for incompatible trials were obtained relative to a neutral trial condition. The LRP revealed a sequential pattern of motor activation. A partial activation of the response corresponding to the prime was followed by a reverse activation pattern. It is argued that these effects primarily reflect an inhibition of the response initially triggered by the prime.

On processing BEASTS and BIRDS: an event-related potential study on the representation of taxonomic structure.

In two event-related potential (ERP) studies it was investigated whether concrete nouns of different taxonomic level (basic level terms, e.g., BIRD, and superordinate terms, e.g., BEAST) are processed differentially under conditions where no processing of taxonomic relations is required. The first experiment, a lexical decision task with taxonomic terms serving as no-go trials, revealed no differences in ERP waveforms to basic level as compared to superordinate terms. The second experiment, a modified oddball paradigm, revealed a consistent pattern of ERP differences: Superordinate deviants elicited a prolonged early positivity as compared to standard items, possibly a P3a component, while basic level deviants elicited an enlarged N400 compared to standard items. Additionally, taxonomic items presented only once in a recognition test elicited different ERPs depending on their respective taxonomic level. Moreover, study- as well as recognition-ERP patterns differed depending on whether Subjects were confronted with basic level terms or with superordinates as standards. It was concluded that representations of basic level terms form a distinct conceptual class, while representations of superordinate terms probably do not.

Explicit and implicit learning of event sequences: evidence from event-related brain potentials.

Event-related brain potentials (ERPs) were recorded during a serial reaction time (RT) task, where single deviant items seldom (Experiment 1) or frequently (Experiment 2) replaced 1 item of a repeatedly presented 10-item standard sequence. Acquisition of sequence knowledge was reflected in faster RTs for standard as compared with deviant items and in an enhanced negativity (N2 component) of the ERP for deviant items. Effects were larger for participants showing explicit knowledge in their verbal reports and in a recognition test. The lateralized readiness potential indicated that correct responses were activated with shorter latencies after training. For deviant items, participants with explicit knowledge showed an initial activation of the incorrect but expected response. These findings suggest that the acquisition of explicit and implicit knowledge is reflected in different electrophysiological correlates and that sequence learning may involve the anticipatory preparation of responses.